Formicoxenus nitidulus

AntWiki: The Ants --- Online
Formicoxenus nitidulus
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Crematogastrini
Genus: Formicoxenus
Species: F. nitidulus
Binomial name
Formicoxenus nitidulus
(Nylander, 1846)

Formicoxenus nitidulus casent0173159 profile 1.jpg

Formicoxenus nitidulus casent0173159 dorsal 1.jpg

Specimen Label


A xenobiont that is unique within Formicoxenus in being able to survive in the nests of at least 11 different ant host species.

At a Glance • Xenobiotic  • Ergatoid queen  


Reddish yellow to brown: whole surface of body smooth and shining with scattered acute pale hairs. Antennal club 3 segmented as long as rest of funiculus: propodeal spines short, set horizontally. Length: 2.8-3.4 mm (Collingwood 1979).

Seifert (2023) - Worker. Numeric data given are arithmetic means based on measurement of 10 specimens (for standard deviation, minimum and maximum values see Tab. 1). Rather small size, CS 642 µm . Head much elongated, CL/CW 1.270. Clypeus extended caudad, semiglobular, in dorsal view with a semicircular anterior margin. Posterior margin of head straight. Frons extremely broad (FRS/CS 0.513). Scape rather short, SL/CS 0.677. Eye small, EYE/CS 0.186, with long setae (Fig. 9). Dorsal profile of promesonotum and propodeum weakly convex, metanotal depression shallow (MGr/CS 1.34 %). Spines rather short (SP/CS 0.143) and not very acute, their bases rather distant (SPBA/CS 0.324). Petiole with strong anterolateral corners, in profile without peduncle and with a broad subpetiolar lobe. Postpetiolar sternite with a well-developed spine. All surfaces of head, mesosoma, waist and appendages smooth and very shiny, sometimes frontal lobes and genae weakly longitudinally carinulate. Gaster glabrous and with very dilute pubescence. Head, mesosoma, waist, scape, femora and tibiae with scattered and short (PnHL/CS 0.077) setae, tapering apicad and round in cross-section. Color variable from light yellowish brown to nearly blackish brown.

As a combination of smooth and shiny body surfaces, very broad frons, elongated head and extended semiglobular clypeus not to confuse.

Keys including this Species


Spain to Eastern Siberia. North Italy to latitude 70º N (Collingwood 1979).

Eurosiberian, temperate-boreal. The very wide host spectrum allows continuous distribution from Scotland and France to Russian Far East. In Fennoscandia up to 70°N, in the Alps ascending to 2300 m. (Seifert, 2023)

Latitudinal Distribution Pattern

Latitudinal Range: 70.377854° to 40.133333°.

Tropical South

Distribution based on Regional Taxon Lists

Palaearctic Region: Austria, Belarus, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland (type locality), France, Germany, Greece, Hungary, Iberian Peninsula, Italy, Latvia, Lithuania, Luxembourg, Netherlands, Norway, Poland, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Türkiye, Ukraine, United Kingdom of Great Britain and Northern Ireland.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.


Collingwood (1979) - This ant occurs only in the nests of Formica rufa and allied mound building species. It is ignored by its host among which the xenobiont species moves freely. Individual nests contain only a few individuals, up to about 100, but often several nests are present within one mound of the host. Nests are located in fragments of wood, hollow twigs, bases of old bracken stems and in the earth floor of the Formica mound. Individuals normally remain concealed within the nests but may wander on the mound surface on warm dull days. It is not known to feed on the Formica brood but in captivity will destroy Leptothorax larvae. Males and winged females may be found during July and August. Mating occurring on the surface of the Formica mound.

Ergatoid queens (i.e. wingless at emergence) are frequent. Mated egg-layers include dealate queens and ergatoid queens. Workers and sexuals are able to solicit food from wood ants, either directly, or the guest ant climbs up the legs and thorax to the head of a host ant that is engaged in food exchange with another Formica, then steals a little food from the droplet the two ants have between their mandibles.

Known Hosts

Formicoxenus nitidulus is known to occur in the nests of the following species (Wilson 1971; Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007).

Flight Period

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


Life History Traits

  • Queen number: monogynous (Buschinger & Winter, 1976; Frumhoff & Ward, 1992) (functional monogyny)


Francoeur et al. (1985) described ergatoid queens to have 1-3 ocelli and other traits resembling more or less the winged queens


Images from AntWeb

Formicoxenus nitidulus casent0173160 head 1.jpgFormicoxenus nitidulus casent0173160 profile 1.jpgFormicoxenus nitidulus casent0173160 profile 2.jpgFormicoxenus nitidulus casent0173160 dorsal 1.jpgFormicoxenus nitidulus casent0173160 label 1.jpg
Queen (alate/dealate). Specimen code casent0173160. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


Images from AntWeb

Formicoxenus nitidulus casent0178346 head 1.jpgFormicoxenus nitidulus casent0178346 profile 2.jpgFormicoxenus nitidulus casent0178346 dorsal 1.jpgFormicoxenus nitidulus casent0178346 label 1.jpg
Male (alate). Specimen code casent0178346. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • nitidulus. Myrmica nitidula Nylander, 1846b: 1058 (w.) FINLAND.
    • Type-material: holotype worker.
    • Type-locality: Finland: Helsingfors (= Helsinki) (F.W. Maeklin).
    • Type-depository: ZMHF.
    • Nylander, 1849: 34 (q.); Mayr, 1855: 419 (m.).
    • Combination in Formicoxenus: Mayr, 1855: 418.
    • Junior synonym of Stenamma westwoodii: Smith, F. 1871b: 4; André, 1874: 203 (in list); Forel, 1874: 102 (in list); Emery & Forel, 1879: 456.
    • Status as species: Nylander, 1849: 34; Foerster, 1850a: 55; Mayr, 1855: 418 (redescription); Nylander, 1856b: 94; Smith, F. 1857b: 40; Smith, F. 1858b: 121; Meinert, 1861: 328; André, 1882c: 273 (in key); André, 1885: 844; Forel, 1886f: 131; Nasonov, 1889: 28; Lameere, 1892: 66; Dalla Torre, 1893: 62; Wasmann, 1894: 162; Emery, 1895c: 271; Ruzsky, 1896: 71; Saunders, E. 1896: 31; Wheeler, W.M. 1900c: 48; Wheeler, W.M. 1901b: 536; Ruzsky, 1902d: 20; Ruzsky, 1902e: 16; Ruzsky, 1905b: 551; Viehmeyer, 1906: 56; Wasmann, 1906: 120; Emery, 1908f: 551; Bondroit, 1910: 499; Stitz, 1914: 66; Forel, 1915d: 18 (in key); Donisthorpe, 1915d: 83; Emery, 1916b: 190; Ruzsky, 1916: 6; Escherich, 1917: 325; Bondroit, 1918: 141; Müller, 1921: 48; Soudek, 1922: 31; Müller, 1923b: 99; Emery, 1924d: 265; Ruzsky, 1925a: 289; Ruzsky, 1925b: 45; Betrem, 1926: 218; Stärcke, 1926: 84 (in key); Donisthorpe, 1927b: 89; Karavaiev, 1927a: 293; Karavaiev, 1927c: 268 (in key); Lomnicki, 1928: 5; Kuznetsov-Ugamsky, 1929a: 30; Kuznetsov-Ugamsky, 1929b: 43; Gösswald, 1932: 96; Arnol'di, 1933b: 598 (in key); Karavaiev, 1934: 147 (redescription); Grandi, 1935: 101; Ruzsky, 1936: 93; Holgersen, 1940: 183; Novák & Sadil, 1941: 84 (in key); Holgersen, 1942: 7; Holgersen, 1944: 168; Ruzsky, 1946: 70; van Boven, 1947: 172 (in key); Dlussky, 1963: 191 (in key); Bernard, 1967: 225 (redescription); Kutter, 1968a: 60; Collingwood & Yarrow, 1969: 73; Baroni Urbani, 1971c: 132; Collingwood, 1971: 160; Bolton & Collingwood, 1975: 3 (in key); Pisarski, 1975: 24; van Boven, 1977: 85; Kutter, 1977c: 144; Arnol’di & Dlussky, 1978: 544 (in key); Collingwood, 1978: 76 (in key); Collingwood, 1979: 78; Francoeur, Loiselle & Buschinger, 1985: 380 (redescription); Agosti & Collingwood, 1987a: 55; Agosti & Collingwood, 1987b: 266 (in key); Nilsson & Douwes, 1987: 58; Radchenko, 1994b: 111 (in key); Bolton. 1995b: 207; Douwes, 1995: 90; Poldi, et al. 1995: 5; Espadaler, 1997b: 29; Gallé, et al. 1998: 215; Czechowski, et al. 2002: 39; Markó & Csösz, 2002: 115; Schulz & Sanetra, 2002: 159; Csösz, & Markó, 2005: 239; Bračko, 2006: 136; Markó, Sipos, et al. 2006: 69; Petrov, 2006: 85 (in key); Bračko, 2007: 17; Seifert, 2007: 230; Radchenko, 2007: 32; Werner & Wiezik, 2007: 140; Zryanin & Zryanina, 2007: 231; Casevitz-Weulersse & Galkowski, 2009: 488; Lapeva-Gjonova, et al. 2010: 19; Boer, 2010: 47; Csösz, et al. 2011: 57; Legakis, 2011: 16; Borowiec, L. & Salata, 2012: 497; Czechowski, et al. 2012: 125; Kiran & Karaman, 2012: 19; Borowiec, L. 2014: 80; Lebas, et al. 2016: 278; Radchenko, 2016: 200; Steiner, et al. 2017: 10; Salata & Borowiec, 2018c: 45; Seifert, 2018: 198; Werner, et al. 2018: 7.
    • Senior synonym of laeviuscula: Mayr, 1855: 418; Nylander, 1856b: 95; Smith, F. 1858b: 121; Dalla Torre, 1893: 63; Ruzsky, 1905b: 551; Donisthorpe, 1915d: 83; Emery, 1924d: 264; Donisthorpe, 1927b: 89; Karavaiev, 1934: 147; Francoeur, Loiselle & Buschinger, 1985: 380; Bolton, 1995b: 207; Radchenko, 2016: 200.
    • Senior synonym of picea: Francoeur, Loiselle & Buschinger, 1985: 380; Bolton, 1995b: 207; Radchenko, 2016: 200.
    • Material of the nomen nudum Myrmica lucidula Smith, F. 1857b: 39 referred here by Donisthorpe, 1915d: 83; Emery, 1924d: 265; Donisthorpe, 1927b: 89; Francoeur, Loiselle & Buschinger, 1985: 380; Bolton, 1995b: 207.
    • Distribution: Austria, Belarus, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Great Britain, Greece, Hungary, Italy, Latvia, Lithuania, Luxembourg, Moldova, Netherlands, Norway, Poland Romania, Russia, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine.
    • [Note: distribution from Borowiec, L. 2014: 80.]
  • laeviuscula. Myrmica laeviuscula Foerster, 1850a: 54 (q.) GERMANY.
    • Type-material: holotype queen.
    • Type-locality: Germany: Aachen (A. Foerster).
    • Type-depository: MNHU.
    • [Note: depository according to Horn & Kahle, 1935: 78.]
    • [Misspelled as laeviuscala by Francoeur, Loiselle & Buschinger, 1985: 380.]
    • Status as species: Schenck, 1852: 132.
    • Junior synonym of Stenamma westwoodii: André, 1874: 203 (in list); Forel, 1874: 102 (in list); Emery & Forel, 1879: 456.
    • Junior synonym of nitidulus: Mayr, 1855: 418; Nylander, 1856b: 95; Smith, F. 1858b: 121; Dalla Torre, 1893: 63; Ruzsky, 1905b: 551; Donisthorpe, 1915d: 83; Emery, 1924d: 264; Donisthorpe, 1927b: 89; Karavaiev, 1934: 147; Francoeur, Loiselle & Buschinger, 1985: 380; Bolton, 1995b: 207; Radchenko, 2016: 200.
  • picea. Leptothorax nitidulus var. picea Wasmann, 1906: 120 (w.) LUXEMBURG.
    • Type-material: syntype workers (number not stated).
    • Type-locality: Luxemburg: (no further data), in nest of Formica pratensis (E. Wasmann).
    • Type-depository: NMLL.
    • Subspecies of nitidulus: Emery, 1908f: 552; Bondroit, 1918: 141; Emery, 1924d: 265; Stärcke, 1926: 85 (in key); Ruzsky, 1946: 70.
    • Junior synonym of nitidulus: Francoeur, Loiselle & Buschinger, 1985: 380; Bolton, 1995b: 207; Radchenko, 2016: 200.

Type Material

Seifert (2023) reports the following:

Myrmica nitidula: This species has been described from an ergatoid male from Helsingfors (=Helsinki), leg F.W. Maeklin. Investigated were two syntypes on different pins. The first is labelled ‘H: fors’, ‘W. Nyland.’, ‘Mus. Fenn’, ‘Mus. Zool. H: fors Spec. typ. No. 5072 Myrmica nitidula Nyl.’ and the second one with equal labels except for ‘... Spec. typ. No. 5073...’.

Myrmica laeviuscula: This taxon has been described from Aachen from a gyne. Types are probably lost but the original description clearly indicates a synonymy with F. nitidulus. Foerster already considered a possible synonymy but decided to describe a new species because his gyne had 11 antennal segments whereas Nylander’s ‘worker’ (in fact an ergatoid male!) was reported to have 12 segments.

Formicoxenus nitidulus var. picea: This taxon was described from Luxembourg as a very dark (nearly blackish brown) color variant found in a nest of Formica pratensis. The synonymy is obvious.


Seifert (2023) redescribed this species using detailed morphometric data (see Table 1 of his study), as explained in the diagnosis of F. nitidulus (see the identification section on this species page) in comparison with other Palaearctic Leptothorax and Formicoxenus.


  • n = 15 (France) (Buschinger et al., 1980; Francoeur et al., 1985; Fischer, 1987).


References based on Global Ant Biodiversity Informatics

  • Agosti, D. and C.A. Collingwood. 1987. A provisional list of the Balkan ants (Hym. Formicidae) and a key to the worker caste. I. Synonymic list. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 60: 51-62
  • AntArea. Accessed on February 5th 2014 at
  • Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
  • Antarea (at on June 11th 2017)
  • ArtDatabanken Bugs (via GBIG)
  • Assing V. 1989. Die Ameisenfauna (Hym.: Formicidae) nordwestdeutscher Calluna-Heiden. Drosera 89: 49-62.
  • Baroni Urbani C., and C. A. Collingwood. 1976. A Numerical Analysis of the Distribution of British Formicidae (Hymenoptera, Aculeata). Verhandlungen der Naturforschenden Gesellschaft in Basel 85: 51-91.
  • Baroni Urbani C., and C. A. Collingwood. 1977. The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. Acta Zoologica Fennica 152: 1-34.
  • Barrett K. E. 1967. Ants in South Brittany. Entomologist's Record and Journal of Variation 79:112-116.
  • Barrett K. E. J. 1968b. The distribution of ants in central southern England. Transactions of the Society for British Entomology 17: 235-250.
  • Behr D., and K. Colln. 1993. Zur ameisenfauna (Hymenoptera, Formicidae) von Gonnersdorf (Kr. Daun). Dendrocopos 20: 148-160.
  • Bernard F. 1967. Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale. Paris: Masson, 411 pp.
  • Bezdecka P. 1996. The ants of Slovakia (Hymenoptera: Formicidae). Entomofauna carpathica 8: 108-114.
  • Blacker N. C. and C. A. Collingwood. 2002. Some significant new records of ants (Hymenoptera: Formicidae) from the Salisbury area, south Wiltshire, England, with a key to the British species of Lasius. British Journal of Entomology and Natural History 15: 25-46
  • Blatrix R., C. Lebas, C. Galkowski, P. Wegnez, P. Pimenta, and D. Morichon. 2016. Vegetation cover and elevation drive diversity and composition of ant communities (Hymenoptera: Formicidae) in a Mediterranean ecosystem. – Myrmecological News 22: 119-127.
  • Blinov V. V. 1984. New for the fauna of Byelorussia ant species. Vestsi Akademii Navuk BSSR. Seryia Biialahichnykh Navuk 1984(5): 113-115.
  • Boer P. 2019. Species list of the Netherlands. Accessed on January 22 2019 at
  • Boer P., W. Dekoninck, A. J. Van Loon, and F. Vankerkhoven. 2003. Lijst van mieren (Hymenoptera: Formicidae) van Belgie en Nederland, hun Nederlandse namen en hun voorkomen. Entomologische Berichten (Amsterdam) 63: 54-58.
  • Boer P., W. Dekoninck, A. J. van Loon, and F. Vankerkhoven. 2003. Lijst van mieren (Hymenoptera: Formicidae) van Belgie en Nederland, hun Nederlandse namen en hun voorkomen. Entomologische Berichten 63(3): 54-57.
  • Boer P., W. Dekoninck, A. J. van Loon, and F. Vankerkhoven. 2003. List of ants (Hymenoptera: Formicidae) of Belgium and The Netherlands, their status and Dutch vernacular names. Entomologische Berichten 63 (3): 54-58.
  • Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
  • Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
  • Borowiec M. L. 2009. New data on the occurrence of formicoxenine ants (Hymenoptera: Formicidae) in Poland. Wiadomo?ci Entomologiczne 28: 237-246.
  • Borowiec M. L. 2009. Nowe dane o rozmieszczeniu mrówek (Hymenoptera: Formicidae) z plemienia Formicoxenini w Polsce. Wiadomosci Entomologiczne 28: 237-246.
  • Boven J. K. A. 1947. Liste de détermination des principales espèces de fourmis belges (Hymenoptera Formicidae). Bulletin et Annales de la Société Entomologique de Belgique 83: 163-190.
  • Boven J. van 1949. Notes sur la faune des Hautes-Fagnes en Belgique. Bulletin et Annales de la Société Entomologique de Belgique 85: 135-143.
  • Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
  • Bracko, G. 2006. Review of the ant fauna (Hymenoptera:Formicidae) of Croatia. Acta Entomologica Slovenica 14(2): 131-156.
  • Casevitz-Weulersse J., and M. Prost. 1991. Fourmis de la Côte-d'Or présentes dans les collections du Muséum d'Histoire Naturelle de Dijon. Bulletin Scientifique de Bourgogne 44: 53-72.
  • Collingwood C. A. 1961. Ants in the Scottish Highlands. Scotish Naturalist 70: 12-21.
  • Collingwood C. A. 1971. A synopsis of the Formicidae of north Europe. Entomologist 104: 150-176
  • Collingwood C.A. 1959. Ants in the Scottish Highlands. The Scottish Naturalist. 70: 12-21
  • Collingwood C.A. 1959. Scandinavian Ants. Entomol. Rec. 71: 78-83
  • Collingwood C.A. 1961. New Vice-County Records for British Ants. Entomologist. 73: 90-93
  • Collingwood C.A. and Satchell J.E. 1956. The Ants of the South Lake District. Journal of the Society for British Entomology. 5: 159-164
  • Collingwood, C. A. 1958b. A key to the species of ants (Hymenoptera, Formicidae) found in Britain. Trans. Soc. Br. Entomol. 13: 69-96
  • Collingwood, C. A. 1964. The Identification of British Ants (Hym. Formicidae). Transactions of the Society for British Entomology. 16:93-121.
  • Collingwood, C. A. 1974. A revised list of Norwegian ants (Hymenoptera: Formicidae). Norsk Entomologisk Tidsskrift 21: 31-35.
  • Collingwood, C. A., and I. H. H. Yarrow. "A survey of Iberian Formicidae." EOS (Revista española de entomología) 44 (1969): 53-101.
  • Collingwood, C. A.. "The Formicidae (Hymenoptera) of Fennoscandia and Denmark." Fauna Entomologica Scandinavica 8 (1979): 1-174.
  • Consani M. 1947. Reperti corologici sulle formiche italiane. Redia 32: 179-182.
  • Csősz S., B. Markó, and L. Gallé. 2011. The myrmecofauna (Hymenoptera: Formicidae) of Hungary: an updated checklist. North-Western Journal of Zoology 7: 55-62.
  • Czechowski W., A. Radchenko, W. Czechowska and K. Vepsäläinen. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw: Natura Optima Dux Foundation, 1-496 pp
  • Czechowski W., and W. Czechowska. 1999. New sites in Poland and notes on the biology of socially parasitic ants Formicoxenus nitidulus (Nyl.) and Harpagoxenus sublaevis (Nyl.) (Hymenoptera, Formicidae). Fragmenta Faunistica 42: 1-6.
  • Dauber, J., J. Bengtsson and L. Lenoir. 2006. Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants. Conservation Biology 20(4):1150-1160
  • Dekoninck W., T. Parmentier, J. Casteels, P. Wegnez, and F. Vankerhoven. 2015. Recente waarnemingen van de glanzende gastmier Formicoxenus nitidulus (Nylander, 1846) in Vlaanderen (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 151: 15-17.
  • Dubovikoff D. A., and Z. M. Yusupov. 2018. Family Formicidae - Ants. In Belokobylskij S. A. and A. S. Lelej: Annotated catalogue of the Hymenoptera of Russia. Proceedingss of the Zoological Institute of the Russian Academy of Sciences 6: 197-210.
  • Dvorak, L., P. BOGUSCH, I. MALENOVSKÝ, P. BEZDÌÈKA, K. BEZDÌÈKOVÁ, K. HOLÝ, P. LIŠKA, J. MACEK, L. ROLLER, M. RÍHA et al. "Hymenoptera of Hády Hill, near the city of Brno (Czech Republic), collected during the Third Czech-Slovak Hymenoptera meeting." Acta Musei Moraviae, Scientiae biologicae (Brno) 93 (2008): 53-92.
  • Else G., B. Bolton, and G. Broad. 2016. Checklist of British and Irish Hymenoptera - aculeates (Apoidea, Chrysidoidea and Vespoidea). Biodiversity Data Journal 4: e8050. doi: 10.3897/BDJ.4.e8050
  • Espadaler X., X. Roig, K. Gómez, and F. García. 2011. Formigues de les Planes de Son i mata de València (Hymenoptera, Formicidae) Treballs de la Institució Catalana d'Història Natural 16: 609-627.
  • Fowles, A.P. 1996. A provisional checklist of the invertebrates recorded from Wales. 2. Aculeate wasps, bees and ants (Hymenoptera: Aculeata). Countryside Council for Wales
  • Francoeur, A., R. Loiselle, and A. Buschinger. "Biosystématique de la tribu Leptothoracini (Formicidae, Hymenoptera). 1. Le genre Formicoxenus dans la région holarctique." Naturaliste Canadien (Québec) 112 (1985): 343-403.
  • Francois J. 1958. Contribution a l'etude ecologique des Formicides (Insectes, Hymenopteres) de la region Dijonnaise. Travaux du laboratoire de Zoologie et de la Station Aquicole Grimaldi de la Faculte des Sciences de Dijon 25, 39 pages.
  • Garcia Garcia F., and A. D. Cuesta-Esgura. 2017. First catalogue of the ants of Burgos province, Spain (Hymenoptera: Formicidae). Boletín de la Sociedad Entomológica Aragonesa 60: 245–258.
  • Grandi G. 1935. Contributi alla conoscenza degli Imenotteri Aculeati. XV. Boll. R. Ist. Entomol. Univ. Studi Bologna 8: 27-121.
  • Holgersen H. 1940. Myrmekologiske notiser I. Nor. Entomol. Tidsskr. 5: 183-187.
  • Holgersen H. 1942. Ants of northern Norway (Hym., Form.). Tromso Mus. Årsh. 63(2): 1-34.
  • Holgersen H. 1943. Ant studies in Rogaland (south-western Norway). Avhandlingar utgitt av det Norske Videnskaps-Akademi i Oslo. I. Matematisk-Naturvidenskapelig Klasse 1943(7): 1-75.
  • Holgersen H. 1944. The ants of Norway (Hymenoptera, Formicidae). Nytt Magasin for Naturvidenskapene 84: 165-203.
  • IZIKO South Africa Museum Collection
  • Jeffery H. G. 1931. The Formicidae (or ants) of the Isle of Wight. Proceedings of the Isle of Wight Natural History and Archaeological Society 2: 125-128. 
  • Karavaiev V. 1927. Übersicht der Ameisenfauna der Krim nebst einigen Neubeschreibungen. Konowia 5: 281-303.
  • Kofler A. 1995. Nachtrag zur Ameisenfauna Osttirols (Tirol, Österreich) (Hymenoptera: Formicidae). Myrmecologische Nachrichten 1: 14-25.
  • Kupianskaia A.N. 1990. Murav'I (Hymenoptera, Formicidae) Dal'nego Vostoka SSSR (1989). Vladivostok. 258 pages.
  • Kvamme T. 1982. Atlas of the Formicidae of Norway (Hymenoptera: Aculeata). Insecta Norvegiae 2: 1-56.
  • Lapeva-Gjonova, L., V. Antonova, A. G. Radchenko, and M. Atanasova. "Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria." ZooKeys 62 (2010): 1-124.
  • Lebas C., C. Galkowski, P. Wegnez, X. Espadaler, and R. Blatrix. 2015. The exceptional diversity of ants on mount Coronat (Pyrénées-Orientales), and Temnothorax gredosi(Hymenoptera, Formicidae) new to France. R.A.R.E., T. XXIV (1): 24 – 33
  • Lelej A. S. 2012. Annotated catalogue of the Insects of Russian Far East. Volume 1. Hymenoptera. Dalnauka: Vladivostok. 635 p.
  • Lomnicki J. 1928. Spis mrówek Lwowa i okolicy. Ksiegi Pamiatkowej (Lecia Gimn. IV Jana Dlugosza Lwowie) 50: 1-10.
  • Maavara V. 1953. Ants of Estonian SSR. ABIKS loodusevaatlejale 10: 1-44.
  • Marikovsky P. I. 1979. Ants of the Semireche Desert. [In Russian.]. Alma Ata: Nauka, 263 pp.
  • Markó B., B. Sipos, S. Csősz, K. Kiss, I. Boros, and L. Gallé. 2006. A comprehensive list of the ants of Romania (Hymenoptera: Formicidae). Myrmecologische Nachrichten 9: 65-76.
  • Markó B., S. Csősz. 2001: Nine new ant species in the Romanian fauna (Hymenoptera: Formicidae): morphology, biology and distribution. – Entomologica Romanica 6: 127-132.
  • Martin, S.J., E.A. Jenner and F.P. Drijfhout. 2007. Chemical Deterrent Enables a Socially Parasitic Ant to Invade Multiple Hosts. Proceedings: Biological Sciences 274(1626):2717-2721
  • Martínez Ibáñez, M. D.. "Algunos datos sobre malformaciones de hormigas." Actas de las VIII Jornadas de la Asociación Española de Entomología (1986): 1035-1041.
  • Müller G. 1921. Due nuove formiche della regione Adriatica. Boll. Soc. Adriat. Sci. Nat. Trieste 27(2): 46-49.
  • Müller, G.. "Le formiche della Venezia Guilia e della Dalmazia." Bollettino della Società Adriatica di Scienze Naturali in Trieste 28 (1923): 11-180.
  • Neumeyer R., and B. Seifert. 2005. Commented check list of free living ant (Hymenoptera: Formicidae) species of Switzerland. Bulletin de la Societe Entomologique Suisse 78: 1-17.
  • Nielsen M. G. 2011. A check list of Danish ants and proposed common names. Ent. Meddr. 79: 13-18.
  • Nylander, W.. "Synopsis des Formicides de France et d'Algérie." Annales des Sciences Naturelles, Zoologie (4)5 (1856): 51-109.
  • Petal J. M. 1963. Faune des fourmis de la reserve de tourbiere en projet a Rakowskie Bagno pres de Frampol (voivodie de Lublin). Annales Universitatis Mariae Curie-Sk?odowska 58(7): 143-174.
  • Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
  • Pisarski B. 1982. Ants (Hymenoptera: Formicoidea) of Warsaw and Mazovia. Memorabilia Zool. 36: 73-90.
  • Punttila P., Y. Haila, and H. Tukia. 1996. Ant communities in taiga clearcuts: habitat effects and species interactions. Ecography 19: 16-28.
  • Radchenko A. G. 2007. The ants (Hymenoptera, Formicidae) in the collection of William Nylander. Fragmenta Faunistica (Warsaw) 50: 27-41.
  • Ruzsky M. 1916. On zoological research in Yeniseisk province, work of summer of 1915. Izv. Imp. Tomsk. Univ. 65 (3rd p part: 1-21.
  • Rzeszowski K., H. Babik, W. Czechowski, and B. Marko. 2013. Ants (Hymenoptera: Formicidae) of Chelmowa Gora in the Swietokrzyski National Park. Fragmenta Faunistica 56(1): 1-15.
  • Saaristo M. I. 1995. Distribution maps of the outdoor myrmicid ants (Hymenoptera, Formicidae) of Finland, with notes on their taxonomy and ecology. Entomol. Fennica 6: 153-162.
  • Schlick-Steiner B. C., and F. M. Steiner. 1999. Faunistisch-ökologische Untersuchungen an den freilebenden Ameisen (Hymenoptera: Formicidae) Wiens. Myrmecologische Nachrichten 3: 9-53.
  • Seifert B. 1994. Die freilebenden Ameisenarten Deutschlands (Hymenoptera: Formicidae) und Angaben zu deren Taxonomie und Verbreitung. Abhandlungen und Berichte des Naturkundemuseums Görlitz 67(3): 1-44.
  • Seifert B. 1998. Rote Liste der Ameisen. - in: M. Binot, R. Bless, P. Boye, H. Gruttke und P. Pretscher: Rote Liste gefährdeter Tiere Deutschlands. Bonn-Bad Godesberg 1998: 130-133.
  • Sellier Y., C. Galkowski, C. Lebas, and P. Wegnez. 2016. Découverte de Temnothorax pardoi (Tinaut, 1987) dans la réserve naturelle nationale du Pinail (Hymenoptera, Formicidae). Revue de l’Association Roussillonnaise d’Entomologie 25(2): 106-113.
  • Sonnenburg H. 2005. Die Ameisenfauna (Hymenoptera: Formicidae) Niedersachsens und Bremens. Braunschweiger Naturkundliche Schriften 7: 377-441.
  • Stroo A. 2003. Het ruggengraatloze soortnbeleid. Nieuwsbrief European Invertebrate Survey – Nederland 36: 8-14.
  • Stumper R. 1953. Etudes myrmecologiques. XI. Fourmis luxembourgeoises. Bulletin Soc. Nat. luxemb. 57: 122-135.
  • Tausan I., M. M. Jerpel, I. R. Puscasu, C. Sadeanu, R. E. Brutatu, L. A. Radutiu, and V. Giurescu. 2012. Ant fauna (Hymenoptera: Formicidae) of Sibiu County (Transylvania, Romania). Brukenthal. Acta Musei 7(3): 499-520.
  • Vespalainen K., B. Pisarski, R. Kantorek, and K. J. Laine. 1984. Formicidae (Hymenoptera) of Inari Lapland. Kevo Notes 7: 115-116.
  • Vogrin, V.. "Prilog fauni Hymenoptera - Aculeata Jugoslavije." Zast. Bilja 31(suppl.) (1955): 1-74.
  • Wegnez P. 2018. Premières decouvertes de Myrmica bibikoffi Kutter, 1963 et de Ponera testacea Emery, 1895, au Luxembourg (Hymenoptera: Formicidae)Bulletin de la Société royale belge d’Entomologie 154: 263–272.
  • Wegnez P., D. Ignace, E. Lommelen, M. Hardy, J. Bogaert, and C. Nilsson. 2015. Redécouverte de Teleutomyrmex schneideriKutter, 1950 dans les Alpes françaises (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 151: 52-57.
  • Wegnez P., S. De Greef, C. Degache, D. Ignace, and W. Dekoninck. 2011. Observations recentes de la fourmi Formicoxenus nitidulus (NYLANDER, 1846) en Belgique et en France (Hymenoptera Formicidae). Bulletin de la Societe Royale Belge d'Entomologie 20-26.
  • Wegnez P., and F. Mourey. 2016. Formica uralensis Ruzsky, 1895 une espèce encore présente en France mais pour combien de temps ? (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 152: 72-80.
  • Wegnez P., and M. Fichaux. 2015. Liste actualisee des especes de fourmis repertoriees au Grand-Duche de Luxembourg (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 151: 150-165
  • Zhuytszyuan D. 2016. The ants (Hymenoptera, Formicidae) Nizhne-Svirsky reserve and their environmental features. Master's thesis Saint Petersburg State University.
  • Zryanin V. A., and T. A. Zryanina. 2007. New data on the ant fauna Hymenoptera, Formicidae in the middle Volga River Basin. Uspekhi Sovremennoi Biologii 127(2): 226-240.